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The

Resting Potential of the Neuron

Messages

in a neuron develop from


disturbances of the resting potential
In the absence of any outside
disturbance, the membrane maintains
an electrical gradient, also known as
polarization a difference in electrical
charge between the inside and outside
of the cell.

The

neuron inside the membrane has a


slightly negative electrical potential
with respect to the outside, mainly
because of negatively charged proteins
inside the cell
This difference in voltage in a resting
neuron is called the resting potential

Forces

ions

acting on sodium and potassium

If charged ions could flow freely across the


membrane, the membrane would depolarize
However, the membrane is selectively
permeable that is, some chemicals pass
through it more freely than others do
(oxygen, carbon dioxide, urea and water
cross freely)
When the membrane is at rest, the sodium
channels are closed, preventing almost all
sodium flow

Certain kinds of stimulation can the sodium


channels.
When the membrane is at rest, potassium
channels are nearly but not entirely closed, so
potassium flows slowly.
Stimulation opens them more widely also, as it
does for sodium channels
The sodium-potassium pump, a protein complex
repeatedly transport three sodium ions out of the
cell while drawing two potassium ions into it
The sodium-potassium pump is an active
transport that requires energy

As

a result, sodium ions are more than


10 times more concentrated outside the
membrane than inside, and potassium
ions are similarly more concentrated
inside than outside
When the neuron is at rest two forces
act on sodium, both tending to push it
into the cell

First,

consider the electrical gradient


Sodium is positively charged and the
inside of the cell is negatively charged
Opposite electrical charges attract, so
the electrical gradient tends to pull
sodium into the cell

Second,

gradient

consider the concentration

The difference in distribution of ions


across the membrane
Sodium is more concentrated outside
than inside, so just by the laws of
probability, sodium is more likely to
enter the cell than to leave it

Why
The

a resting potential?

body invests much energy to


operate the sodium-potassium pump,
which maintains the resting potential
Why is it worth so much energy?
The resting potential prepares the
neuron to respond rapidly

The

action potential

Messages

sent by axons are called action


potentials
When an axons membrane is at rest, the
recordings show a negative potential
inside the axon
If we now use another electrode to apply
a negative charge, we can further
increase the negative charge inside the
neuron
The charge is called hyperpolarization,
which means increased polarization

When

the stimulation ends, the charge


returns to its original resting level.
Applying a stronger current, stimulation
beyond the threshold of excitation,
produces a massive depolarization of the
membrane
When the potential reaches the
threshold, the membrane opens its
sodium channels and permits sodium ions
to flow into the cell
The potential shoots up far beyond the
strength of the stimulus

depolarization that passes the


threshold produces an action potential
One that falls short of the threshold
does not produce an action potential

At the start, sodium ions are mostly outside


the neuron and potassium ions are mostly
inside
When the membrane is depolarized, sodium
and potassium channels in the membrane
open
At the peak of the action potential, the
sodium channels close

protein that allows sodium to cross is


called sodium channel, one that allows
potassium to cross is a potassium
channel, and so forth
The ones regulating sodium and
potassium are voltage-gated channels
That is, their permeability depends on
the voltage difference across the
membrane

At

the resting potential, the sodium


channels are closed (permitting no
sodium to cross) and the potassium
channels are almost closed (allowing only
a little flow of potassium)
As the membrane becomes depolarized,
both the sodium and the potassium
channels begin to open, allowing freer
flow

When the depolarization reaches the threshold


of the membrane, the sodium channels open
wide enough for sodium to flow freely
Driven by both the concentration gradient and
the electrical gradient, the sodium ions enter
the cell rapidly, until the electrical potential
across the membrane passes beyond zero to a
reversed polarity
Remember that depolarizing the membrane also
opens potassium channels

When

the membrane is at rest, are the


sodium ions more concentrated inside the
cell or outside> where are the potassium ions
more concentrated?

When

the membrane is at rest, what tends to


drive the potassium ions out of the cell?
What tends to draw them into the cell?

The

all-or-none law

the amplitude and velocity of an action


potential are independent of the intensity of
the stimulus that initiated it, provided that
the stimulus reaches the threshold
the law puts constraints on how an axon can
send a message
To signal the difference between a weak
stimulus and a strong stimulus, the axon cant
send bigger or faster action potential
All it can change is the timing

For example, a taste axon shows one rhythm for


sweet tastes and a different rhythm for bitter
tastes

The

refractory period
Immediately after an action potential,
the cell is in a refractory period during
which it resists the production of further
action potentials
In the first part of this period, the
absolute refractory period, the
membrane cannot produce an action
potential, regardless of the stimulation
During the second part, the relative
refractory period, a stronger than usual
stimulus is necessary to initiate an action
potential

The

refractory period has two


mechanism:

The sodium channels are closed, and


potassium is flowing out of the cell at a
faster than usual rate

How

action potential moves down the axon

In a motor neuron, an action potential begins


on the axon hillock, a swelling where the axon
exits the soma
During the action potential, sodium ions enter
a point on the axon
The term propagation of the action potential
describes the transmission of an action
potential down an axon

Lets

review the action potential


When an area of the axon membrane
reaches its threshold of excitation,
sodium channels and potassium channels
open
At first, the opening of potassium
channels produces little effect
Opening sodium channels lets sodium ions
rush into the axon
Positive charge flows down the axon and
opens voltage-gated sodium channels at
the next point

At

the peak of the action potential, the


sodium gates snap shut.
They remain closed for the next
millisecond or so, despite the
depolarization of the membrane

Because

the membrane is
depolarized, voltage-gated
potassium channels are open
Potassium ions flow out of the axon,
returning the membrane toward its
original depolarization
A few milliseconds later, the voltagedependent potassium channels close

The

myelin sheath and saltatory


conduction
In

the thinnest axons, action potentials


travel at a velocity of less than 1 m/s.
To increase the speed still more,
vertebrate axons evolved a special
mechanism: sheaths of myelin, an
insulating material composed of fats and
protein
Myelinated axons, those covered with a
myelin sheath, found only in vertebrates,
are covered with layers of fats and
proteins

The

myelin sheath is interrupted


periodically by short sections of axon
called nodes of Ranvier, each one about 1
micrometer wide.
In most cases, the action potential starts
at the axon hillock, but in some cases it
starts at the first node of Ranvier
The jumping of action potentials from
node to node is referred to as saltatory
conduction,

In

addition to providing rapid conduction


of impulses, saltatory conduction
conserves energy: instead of admitting
sodium ions at every point along the axon
and then having to pump them out via
the sodium-potassium pump, a
myelinated axon admits sodium only at
its nodes
An axon that never had a myelin sheath
conducts impulses slowly but steadily
An axon that has lost its myelin is not the
same

After

myelin forms along an axon, the


axon loses its sodium channels under the
myelin
If the axon later loses its myelin, it still
lacks sodium channels in the areas
previously covered with myelin, and most
action potentials die out between one
node and the next

Local

neurons

Axons

produce action potentials


However, many small neurons have
no axons
Neurons without an axon exchange
information with only their closest
neighbors
We therefore call them local
neurons

Because they do not have an axon, they


do not follow the all-or-none law

When

a local neuron receives information


from other neurons, it has a graded
potential, a membrane potential that
varies in magnitude in proportion to the
intensity of the stimulus
The change in membrane potential is
conducted to adjacent areas of the cell,
in all directions, gradually decaying as it
travels
Those various areas of the cell contact
other neurons, which they excite or
inhibit through synapses

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